Two new species of Endecous Saussure, 1878 (Orthoptera: Grylloidea: Phalangopsidae) from caves in central-western Brazil, with the proposition of a new subgenus to accommodate E. vitreus Bolfarini & Campos, 2023 Author Carvalho, Pedro Henrique Mendes 0000-0003-0859-0255 Centro de Estudos em Biologia Subterrânea (www. biologiasubterranea. com. br), Setor de Biodiversidade Subterrânea, Departamento de Ecologia e Conservação, Universidade Federal de Lavras, Cx Postal 3037, Campus Universitário, CEP 37200 - 000, Lavras, Minas Gerais, Brazil & carvalhopedrohm @ gmail. com; https: // orcid. org / 0000 - 0003 - 0859 - 0255 carvalhopedrohm@gmail.com Author Castro-Souza, Rodrigo Antônio 0000-0002-3439-9991 Laboratório de Macroecologia e Conservação da Biodiversidade, Departamento de Botânica e Ecologia, Instituto de Biociências, Universidade Federal de Mato Grosso, CEP 78060 - 900, Cuiabá, MT, Brazil & rodrigodesouzaac @ gmail. com; https: // orcid. org / 0000 - 0002 - 3439 - 9991 rodrigodesouzaac@gmail.com Author Ferreira, Rodrigo Lopes Centro de Estudos em Biologia Subterrânea (www. biologiasubterranea. com. br), Setor de Biodiversidade Subterrânea, Departamento de Ecologia e Conservação, Universidade Federal de Lavras, Cx Postal 3037, Campus Universitário, CEP 37200 - 000, Lavras, Minas Gerais, Brazil text Zootaxa 2023 2023-10-10 5353 3 201 234 http://dx.doi.org/10.11646/zootaxa.5353.3.1 journal article 10.11646/zootaxa.5353.3.1 1175-5326 8427318 A8B27F7F-5E7E-4DD4-8E92-A52DD0072BD1 Endecous ( Endecous ) liviae n. sp. ( Figures 2–6 , 7–15 , 16–19 , 20–27 , 28–32 , 33–36 , 37–39 , 40–43 ; Table 1 ) Etymology— The specific epithet of the species is dedicated to the biospeleologist Lívia Medeiros Cordeiro Borghezan, whose dedication to the study and preservation of caves in the Serra da Bodoquena karst region has been exceptional. The specific epithet is expressed as a Latinized adjective, paying homage to her significant contributions. Material examined— Holotype , ♂, code ISLA 106153 , Brazil , Mato Grosso do Sul , municipality of Bodoquena , Gruta do Alex II cave ( 56°42’35.06”W , 20°36’19.48”S ), 02.X.2022 , R.L. Ferreira ; condition: head, right tegmen, legs I, II and III detached and stored alongside the holotype , phallic complex dissected and also stored alongside the holotype . Allotype : 1 ♀ , ISLA 106154, same data as the holotype; condition: legs I, II and III detached and stored alongside the alotype, copulatory papilla dissected and also stored alongside the alotype. Paratypes : 1 ♂, ISLA 106152, 1 ♀ , ISLA 106155, same locality and data as the holotype ; 4 ♂♂, ISLA 106132, ISLA 106133, ISLA 106134, ISLA 106135, municipality of Bonito , Gruta América cave ( 56°36’39.01”W , 21°11’28.89”S ), 29.IX.2022 , R . L. Ferreira ; 5 ♂♂, ISLA 106144, ISLA 106145, ISLA 106146, ISLA 106147, ISLA 106148, 3 ♀♀ , ISLA 106149, ISLA 106150, ISLA 106151, municipality of Bonito , Gruta Terra Planetária cave ( 56°36’8.46”W , 21° 8’4.36”S ), 24.IX.2022 , R . L. Ferreira ; 1 ♂, ISLA 106156, 1 ♀ , ISLA 106158, municipality of Bodoquena , Gruta Dona Benedita cave ( 56°43’23.51”W , 20°34’0.55”S ), 26.IX.2022 , R . L. Ferreira ; 1 ♂, ISLA 106159, 1 ♀ , ISLA 106160, municipality of Bodoquena , Gruta Manoel Cardoso cave ( 56°43’23.39”W , 20°34’7.12”S ), 26.IX.2022 , R . L. Ferreira ; 2 ♂♂, ISLA 106171, ISLA 106172, 1 ♀ , ISLA 106173, municipality of Bodoquena , Gruta do Bel I cave ( 56°42’59.17”W , 20°35’20.53”S ), 03.X.2022 , R . L. Ferreira ; 1 ♀ , ISLA 106174, municipality of Bodoquena , Gruta do Bel II cave ( 56°43’0.48”W , 20°35’22.19”S ), 03.X.2022 , R . L. Ferreira ; 1 ♂, ISLA 106175, 1 ♀ , ISLA 106176, municipality of Bonito , Gruta Catedral cave ( 56°50’46.90”W , 20°56’17.52”S ), 05.X.2022 , R . L. Ferreira ; 2 ♂♂, ISLA 106177, ISLA 106178, 2 ♀♀ ISLA 106179, ISLA 106180, municipality of Bonito , Gruta do Lago Azul cave ( 56°35’28.91”W , 21°8’40.40”S ), 19.IX.2004 , R . L. Ferreira ; male paratypes condition: right tegmen and legs I, II and III detached and stored alongside the paratype , phallic complex dissected and also stored alongside the paratype ; female paratypes condition: legs I, II and III detached and stored alongside the paratype , copulatory papilla dissected and also stored alongside the paratype . TABLE 1. Endecous (Endecous) liviae n. sp. , morphological measurements (mm), mean (M) and standard deviation (SD) of 10 male and five female adult specimens; pars stridens refers to the number of teeth of the stridulatory file; # indicates that the right appendage, instead of the left one, was measured; * unable to measure due to damage in the structure.

♂	106133	106134	106145	106146	106152	106153	106156	106159	106171	106172	M	SD
Head width	3.372	3.054	2.887	2.778	2.772	2.733	2.830	2.995	2.827	2.757	2.901	0.196
Head length	2.440	2.313	2.159	2.225	2.007	1.989	2.143	2.295	1.950	2.075	2.160	0.159
Intraocular distance	1.831	1.624	1.578	1.496	1.502	1.552	1.561	1.605	1.533	1.501	1.578	0.099
Compound eye	0.537	0.491	0.451	0.417	0.414	0.367	0.415	0.439	0.403	0.388	0.432	0.050
width
Compound eye	1.005	0.958	0.914	0.879	0.894	0.789	0.866	0.896	0.877	0.826	0.890	0.061
length
Maxillary	1.913	1.810	1.463	1.701	1.494	1.479	1.650	1.473	1.611	1.692	1.629	0.155
palpomere III
Maxillary	1.977	2.074	1.334	1.898	1.844	1.606	1.825	1.868	1.902	1.871	1.820	0.208
palpomere IV
Maxillary	2.571	2.554	1.768	2.360	2.068	1.953	2.251	2.202	2.459	2.276	2.246	0.260
palpomere V
Pronotum width	4.230	4.306	3.688	3.970	3.876	3.817	3.922	4.254	3.947	3.985	4.000	0.202
Pronotum length	2.896	2.619	2.482	2.478	2.522	2.376	2.417	2.610	2.506	2.568	2.547	0.145
Right tegmen width	4.206	3.999	3.810	3.588	3.487	3.588	3.635	3.958	3.619	3.600	3.749	0.234
Right tegmen length	4.927	4.941	4.495	4.435	4.496	4.435	4.395	4.859	4.685	4.582	4.625	0.213
Leg I length	20.878	18.772	17.465	17.181	16.355	15.353#	16.088	17.601	16.536#	16.525#	17.763	1.631
Leg II length	21.614	19.187	18.069	17.876	16.359	15.725#	16.413	17.806	16.839#	16.921	18.031	1.728
Femur III length	12.901	11.794	10.813	10.764	10.440	9.787#	10.080	11.404	10.277#	10.173	11.046	0.951
Tibia III length	13.732	13.110	11.600	11.586	10.962	10.293#	10.763	11.787	10.888#	11.048	11.824	1.061
Body length	23.051	19.805	21.971	19.074	19.833	15.597	17.763	18.107	18.876	17.543	19.162	2.170
Pars stridens	75	72	58	60	62	64	63	66	71	72	66.300	5.832

♀	106149	106151	106154	106173	106158	M	SD
Head width	3.674	3.609	3.126	3.121	3.425	3.391	0.261
Head length	2.632	2.612	2.326	2.386	2.543	2.500	0.137
Intraocular distance	1.957	1.946	1.695	1.606	1.865	1.814	0.156
Compound eye width	0.609	0.526	0.513	0.462	0.537	0.532	0.074
Compound eye length	1.172	1.126	0.852	0.940	1.021	1.022	0.131
Maxillary palpomere III	2.199	2.221	1.652	1.777	1.844	1.939	0.257
Maxillary palpomere IV	2.193	2.587	1.898	2.025	2.118	2.164	0.261
Maxillary palpomere V	2.911	3.162	2.464	2.403	2.804	2.749	0.317
Pronotum width	4.987	4.882	4.527	4.214	4.645	4.651	0.305
Pronotum length	3.397	3.292	2.776	2.635	3.089	3.038	0.327
Femur III length	13.968	14.248	11.584	10.609	12.520	12.586	1.548
Tibia III length	14.854	15.138	12.346	11.525	13.076	13.388	1.570
Body length	24.936	25.694	21.027	18.915	24.040	22.922	2.856
Ovipositor length	13.831	14.262	10.887	10.462	*	12.361	1.962

FIGURES 2–6 . Endecous ( Endecous ) liviae n. sp. (ISLA 106153) phallic complex. (2) dorsal view, (3) ventral view, (4) frontal view, (5) laterofrontal view, (6) lateral view. Abbreviations: A, pseudepiphallic ventral branch (or “A” sclerite); Ps.arm, pseudepiphallic arm; Ps.db, pseudepiphallic dorsal branch; Ps.ib, pseudepiphallic inner bars; R, pseudepiphallic rami; Ps.p, pseudepiphallic paramere; Ect.ap, ectophallic apodeme; Ect.arc, ectophallic arc; Ect.lb, ectophallic lateral bar; Ect.mp, ectophallic median projection; End.sc.a, endophallic sclerite anterior portion; End.sc.d, endophallic sclerite duct; End.sc.p, endophallic sclerite posterior portion. Diagnosis —Combination of the following characteristics: pseudepiphallic dorsal branches bending towards the center of the phallic complex, broadened at the base and tapered towards the apex, which is rounded ( Figs. 2, 4–6 , Ps.db); pseudepiphallic rami elongated and triangular in shape ( Figs. 2–6 , R); inner bars of the ectophallic invagination forming a V-shaped structure, with a less sclerotized, subtriangular in shape, dorsocentral projected extension ( Figs. 2, 4–6 , Ps.ib); ectophallic arc well-developed, conical and rounded, central portion bordering the base of the ectophallic median projections ( Figs. 2 and 3 , Ect.arc); anterior portion of the endophallus well-developed, sclerotized, with a long apodeme on the opposite side of the central groove ( Figs. 2, 3 and 6 , End.sc.a). Male phallic sclerites ( holotype ISLA 106153, Figs.2–6 )—Phallic complex longer than broad and dorsoventrally flattened. Pseudepiphallus : arms slightly elongated and broadened laterally ( Fig. 2 , Ps.arm); dorsal branches sclerotized, bending towards the center of the phallic complex, broadened at the base and tapered towards the apex, which is rounded ( Figs. 2 and 4 , Ps.db); ventral branches elongated and sclerotized, reaching the dorsolateral portion of the pseudepiphallic parameres ( Figs. 5 and 6 , A); paramere 1 and 2 fused in a single sclerotized and concave structure, C-shaped in dorsal view, proximal end ventrally projected, bending and pointing outwards ( Figs. 2 and 3 , Ps.p); inner bars slightly concave, inclined, forming a V-shaped structure, with a less sclerotized, subtriangular in shape, dorsocentral projected extension in dorsal view ( Fig. 2 , Ps.ib); rami elongated, slightly longer than the ectophallic apodeme in dorsal and ventral views, broadened at the base, tapered at the end, triangular in shape in lateral view ( Figs. 2, 3 and 6 , R). Ectophallic invagination : ectophallic arc well-developed, conical and rounded, central portion bordering the base of the ectophallic median projections ( Fig. 3 , Ect.arc); apodemes short and slightly sclerotized, distal end slightly curved inwards ( Figs. 2, 3 and 6 , Ect.ap); median projections thin, sinuous and barely sclerotized, reaching the pseudepiphallic parameres in dorsal view ( Fig. 2 , Ect.mp); lateral bars well-developed, elongated, curved outwards, tapered at the apex, reaching the outer ventral portion of the pseudepiphallic parameres ( Figs. 3, 5 and 6 , Ect.lb). Endophallus : anterior portion well-developed, sclerotized, with a long apodeme on the opposite side of the central groove ( Figs. 2, 3 and 6 , End.sc.a); duct membranous, elongated, longer than the ectophallic apodeme and widened dorsoventrally at the connection to the sclerite anterior portion ( Figs. 2, 3 and 6 , End.sc.d); posterior portion membranous and well-developed, dorsally projected, almost touching the pseudepiphallic dorsal branches ( Figs. 3 and 6 , End.sc.p). Variations in phallic sclerites ( holotype and paratypes , n = 7, ISLA 106132, ISLA 106133, ISLA 106144, ISLA 106145, ISLA 106153, ISLA 106156, ISLA 106159)—Phallic complexes vary slightly in size and degree of sclerotization, being more or less flattened dorsoventrally; pseudepiphallic arms (Ps.arm) vary slightly in length; pseudepiphallic dorsal branches (Ps.db) vary slightly in degree of inclination towards the center of the phallic complex, the tips almost touching or slightly distant from one another, varying slightly in sinuosity; pseudepiphallic inner bars (Ps.ib) vary slightly in degree of inclination, dorsocentral extension varying in size and varying slightly in shape; ectophallic arcs (Ect.arc) central extension vary in length; ectophallic apodemes (Ect.ap) vary slightly in degree of sinuosity and sclerotization; endophallic sclerite posterior portion (End.sc.p) more or less projected dorsally. Morphology ( holotype ISLA 106153, Figs. 7–20 )— Body color : vertex, scape, pedicel and flagellum yellowish brown; fastigium dark brown; front and gena varying from light yellow to white, partially covered by a dotted brown mark; clypeus and labrum also varying from light yellow to white; mandibles and maxiles yellowish brown, galea and labium white; ommatidia black, with a slightly depigmented area near the base of the antennal scapes ( Figs. 9–11 ); maxillary palpomeres I and II whitish, maxillary palpomeres III, IV and V light yellow, palpomere V white at the tip; all labial palpomeres light yellowish brown, labial palpomere III white at the tip ( Figs. 10 and 11 ); pronotum, tegmina and abdomen yellowish brown, tergites distal portion brown ( Figs. 7, 8, 12 and 13 ); supra-anal plate yellowish brown, subgenital plate light yellow proximally, whitish in the central to distal region, and darkened towards the distal end ( Figs. 13, 14 and 15 ); cerci yellowish brown, darker at first sight due to the presence of setae ( Fig. 13 ); legs yellowish brown, white at their base ( Figs. 16–19 ). Head : slightly pubescent, elongated in frontal view; fastigium as a short and broad extension of the vertex, inclined and pointing downwards, with long bristles; mandibles sclerotized at the apex and lateral margins; maxiles sclerotized at the apex; maxillary palpomeres I and II short and same-sized, III–V longer, palpomere V claviform; labial palpomeres I–III increasing in size, palpomere III dilated at the apex; compound eyes reduced and elliptical, ocelli absent ( Figs. 9–11 ). Thorax : pronotum dorsal disk broader than long ( 3.82 mm and 2.38 mm in width and length, respectively), pubescent, anterior and posterior margins arched and covered in long bristles, lateral lobes subtriangular in shape and slightly shifted towards the head ( Figs. 7 and 12 ). Right tegmen : slightly sclerotized; covering the first two urotergites ( 3.59 mm and 4.43 mm in width and length, respectively); harp with one complete and four incomplete crossveins (two bifurcated proximally); mirror subtriangular, with one arched crossvein, and two cells; basal field with two secondary veins connecting Cu2 to 1A, 1A and 2A veins connected through a poorly-marked secondary vein; lateral field with two almost completely parallel longitudinal veins and several anastomotic poorly-marked secondary veins ( Figs. 8, 12 and 20 ); stridulatory file with 64 teeth. Abdomen : cerci pubescent, with long bristles throughout all their extension and globose setae at their base, mostly on the inner side ( Figs. 13–15 ); supra-anal plate shorter than the subgenital plate, subtriangular, distal margin rounded and covered in long bristles, lateral projections short and rounded, paraprocts slightly longer than the supra-anal plate ( Figs. 13 and 14 ); subgenital plate proximal margin substraight, distal margin rounded, with a small dent in the center and covered in bristles ( Figs. 13–15 ). Legs : pubescent. Leg I ( Figs. 16 and 17 ): tibia with an oval tympanum on its inner side and two ventral apical spurs; tarsomere I ventrally serrated and longer than tarsomeres II and III together. Leg II ( Figs. 16 and 17 ): tibia with two ventral apical spurs of equal size and two lateral short spurs, one on each side; tarsomere I ventrally serrated and longer than tarsomeres II and III together. Leg III ( Figs. 18 and 19 ): femur developed; tibia longer than the femur ( 10.29 mm and 9.79 mm , respectively); tibia armed with three subapical spurs on the inner side ( Fig. 19 ; m, n, o), four on the outer side ( Fig. 18 ; w, x, y, z), the most distal one, “z”, being the shortest, four apical spurs on the inner side ( Fig. 19 ; d, e, f, g), spurs “e” and “f” longer the “d” and “g”, and three on the outer side ( Fig. 18 ; a, b, c), spur “a” being the longest and “c” the shortest; tarsomere I longer than tarsomeres II and III together, ventrally serrated, with two apical spurs, the inner one being the longest. FIGURES 7–15 . Endecous ( Endecous ) liviae n. sp. (ISLA 106153) male general morphology. (7) pronotum in dorsal view; (8) right tegmen in dorsal view (9) head in dorsal view; (10) head in frontal view; (11) head and palpi in lateral view, compound eye highlighted; (12) pronotum and tegmen in lateral view; (13) supra-anal and subgenital plates in lateral view; (14) supra-anal plate in dorsal view; (15) subgenital plate in ventral view. FIGURES 16–19 . Endecous ( Endecous ) liviae n. sp. (ISLA 106153) right legs. (16) inner view: leg I, tympanum highlighted, and leg II, apical spurs highlighted; (17) outer view: leg I and leg II, apical spurs highlighted; (18) outer view: leg III, apical spurs highlighted; (19) inner view: leg III, apical spurs highlighted. FIGURES 20–27 . Endecous ( Endecous ) liviae n. sp. right tegmina, dorsal and lateral view. (20) holotype, ISLA 106153; (21) paratype, ISLA 106171; (22) paratype, ISLA 106133; (23) paratype, ISLA 106134; (24) paratype, ISLA 106145; (25) paratype, ISLA 106146, (26) paratype, ISLA 106156, (27) paratype, ISLA 106159. FIGURES 28–32 . Endecous ( Endecous ) liviae n. sp. (ISLA 106154) ovipositor. (28) supra-anal plate; (29) subgenital plate; (30) ovipositor, lateral view; (31) ovipositor apex, dorsal view; (32) ovipositor apex, lateral view. FIGURES 33–36 . Endecous ( Endecous ) liviae n. sp. copulatory papillae. (33) allotype, ISLA 106154; (34) paratype, ISLA 106155; (35) paratype, ISLA 106149; (36) paratype, ISLA 106173; A—dorsal view; B—ventral view; C—lateral view. Variations in right tegmina ( holotype and paratypes , n = 8, ISLA 106133, ISLA 106134, ISLA 106145, ISLA 106146, ISLA 106153, ISLA 106156, ISLA 106159, ISLA 106171, Figs. 20–27 )— Stridulatory file with 65 ± 5.16 teeth (n = 10, holotype and paratypes ). Harp with one ( Figs. 20 and 23 ), two ( Figs. 22, 24 and 26 ), three ( Figs. 25 and 27 ) or four ( Fig. 21 ) complete diagonal crossveins; incomplete crossveins may be present, some of them reaching a light-colored longitudinal striation ( Figs. 20 and 23 ). Mirror with one ( Fig. 20 ), two ( Figs. 21–24, 26 and 27 ) or three ( Fig. 25 ) complete crossveins, the second and third most distal ones are connected through a secondary vein; an incomplete crossvein may be present ( Fig. 27 ); mirror outline inconsistent due to variations in the form of the distal margin. Basal field with poorly-marked secondary veins branching out of 1A and reaching Cu2; 1A may be directly connected to Cu2 ( Fig. 26 ); 1A may also be fragmented in two and connected to the central portion of Cu2 ( Fig. 24 ); 1A, 2A and 3A well-marked. Lateral field with two parallel longitudinal veins that may or may not be connected through short secondary veins; anastomosis of poorly-marked secondary veins branching out of the longitudinal veins; secondary veins reaching the right margin of the tegmina. Female morphology ( allotype ISLA 106154 Figs. 28–32 )—same pattern of body coloration as the male, but slightly darker; bigger than the male (22.92 ± 2.86 mm and 19.16 ± 2.17 mm in length, respectively); apterous; supra-anal plate subtriangular in shape, pubescent, with long bristles covering the distal margin, which is rounded, lateral projections short and rounded ( Fig. 28 ); subgenital plate pubescent, lateral margins angled inwards from the base, distal margin W-shaped ( Fig. 29 ); ovipositor ( 10.89 mm in length) shorter than the cerci, shaped like a curved sword, with a dorsoventral constriction near the tip, apex acute like an arrow-head ( Figs. 30–32 ). Copulatory papilla ( allotype ISLA 106154, Fig. 33 )—sclerotized and chalice-like; anterior potion broadened, ventral side with a less sclerotized V-shaped region, like a deep indentation, ventral margin slightly longer than the dorsal margin; posterior portion membranous with a small circular opening on the ventral side. Variations in copulatory papillae ( allotype and paratypes , ISLA 106149, ISLA 106154, ISLA 106155, ISLA 106173, Figs. 33–36 )—less sclerotized region on the ventral side may be reduced in size and U-shaped; a small Vshaped dent may be present in the center of the dorsal margin. FIGURES 37–39 . Endecous ( Endecous ) liviae n. sp. habitat. (37) Gruta América cave, external environment; (38) Gruta América cave, entrance; (39) Gruta América cave, internal environment. Photo credits: (37) Rodrigo Lopes Ferreira; (38) and (39) Philippe Crochet. FIGURES 40–43 . Endecous ( Endecous ) liviae n. sp. (40) adult male of E. ( E. ) liviae n. sp. , Gruta Alex II cave; (41) adult female of E. ( E. ) liviae n. sp. , Gruta Alex II cave; (42) adult male of E. ( E. ) liviae n. sp. , Gruta Terra Planetária cave; (43) adult female of E. ( E. ) liviae n. sp. , Gruta América. Photo credits: Rodrigo Lopes Ferreira. Ecological remarks —Specimens of Endecous ( E. ) liviae n. sp. were found in nine caves located in the municipalities of Bodoquena (Gruta Dona Benedita, Gruta Manoel Cardoso, Gruta do Bel I, Gruta do Bel II and Gruta Alex II caves) and Bonito (Gruta América— Figs. 38 and 39 , Gruta Catedral, Gruta do Lago Azul and Gruta Terra Planetária caves). Given the limited sampling of caves conducted by the authors, it is probable that the species has a widespread distribution in the region, potentially occurring in additional caves. Unfortunately, the once pristine vegetation in the area has undergone significant alterations over the past few decades due to extensive agricultural and pasture activities. Consequently, the landscape has become highly fragmented, with the remaining forests largely associated with the limestone outcrops ( Fig. 37 ). This inference is supported by the abundance of caves in the area, coupled with the considerable distance between the caves where the three populations were found ( 68 km in a straight line between the farthest caves). Most adult specimens were observed on the caves’ walls ( Figs. 40 and 41 ), while in several locations, they were found concealed within speleothems and rock crevices ( Fig. 43 ). This behavior suggests a certain degree of photophobia, as the presence of researchers equipped with headlamps may disturb them, causing them to seek shelter. Nevertheless, adult specimens, particularly males, could be easily found while engaging in vocalizations as part of their courtship behavior aimed at attracting females ( Fig. 42 ).Another noteworthy observation is that the majority of specimens found in all caves were in their immature stages. Consequently, locating adult individuals proved to be more challenging, potentially indicating the difficulty in reaching adulthood. This can be attributed to the presence of several predators within the caves that prey upon immature specimens. As a result, only a small proportion of the immature individuals are likely to successfully transition into adulthood due to the substantial predation pressure they face.